Odonatologica 4(1): 1-9 March I, 1975
Endemic dragonflies of Late Pleistocene age of the Hula Lake area (northern Israel), with notes on the Calopterygidae of the rivers Jordan (Israel, Jordan) and Litani (The Lebanon), and description of Urothemis
edwardsi hulae subspec. nov. (Libellulidae)*
H.J. Dumont
Institute of Zoology, University of Ghent, Ledeganckstraat 35,
B-9000 Ghent, Belgium
Received January 7, 1975
Volcanic events during the middle Pleistocene blocked the Jordan Rift at
the frontier between Israel and the Lebanon. South of it, in the Hula Lake
and area, four endemic subspecies of dragonflies evolved, presumably during
after the Wiirm. They are all subspecies to Ethiopian species. North of the
barrier evolved Calopteryx hyalina Martin, a taxon of uncertain status. The
two endemic Anisoptera of Lake Hula (Rhyothemis semihyalinasyriaca Selys
and extinct while Urothemis edwardsi hulae ssp.n.) are probably now, the
Zygoptera ( Pseudagrion torridum hulae Dumont and P. sublacteum mortoni
Ris & Schmidt) are still in existence.
INTRODUCTION
The dragonfly fauna of the Mediterranean coast of the Near East is quite
of of complex. In terms present-day zoogeographical divisions, it consists a
of majority Mediterranean elements, with an immixture of Ethiopian and
Oriental species. In addition, a number of endemics occur. It is the nature of
forms some among these endemics that the object of this paper.
* Paper presented at the Fourth Colloquium of Dutch and Belgian Dragonfly Workers,
Utrecht, December 14, 1974,
1 THE ENDEMICS OF THE UPPER JORDAN VALLEY
(LAKE HULA AREA)
PSEUDAGRION TORRIDUM HULAE DUMONT, 1974
Restricted to Lake Hula where it is common. Described and figured in
DUMONT, 1974. Structurally, it is almost identical to P. t. torridum Selys. This is widely distributed in Africa, reaching Egypt and the South-Western edge of the Sinai desert, which is the North-Eastern limit ofits known distribution.The present-time disjunction of the Hula population is thus about 700 km.
E.C.G. Fig. 1 (a) Hind wing of Rhyothemis s. semihyalina, <5, Maun, Botswana (leg. Pinhey);
- (b) Hind wing of R. semihyalinasyriaca, 9, Selys’ type from ’’Syria”.
2 PSEUDAGRION SUBLACTEUM MORTONI (RIS & SCHMIDT, 1936)
in Occurring on Lake Hula but also other places of the Jordan valley (see map in DUMONT, 1974). Present disjunction with the Ethiopian stock unknown, since P. s. sublacteum(Karsch) might occur in Egypt.
RHYOTHEMIS SEMIHYALINA SYRIACA SELYS, 1850
Figures 1 b; 2 a, b
female Described by Selys on a in the Latreille collection, labeled ’’Syria” without any further indication. According to the description, it differs from typical semihyalina in having the black spot at the base of the hindwings more reduced, leaving a narrow hyaline fringe along the posterior border of the wing. I found and these characters well exemplified on the type (Brussels Museum) on the following additional material: 266, Department of Genetics, The Hebrew
University of Jerusalem, one without label, one labeled Hula, 23.VI.1952; 10 specimens in the Department of Zoology, University of Tel Aviv (7 from Hula,
of one from Daphne north Hula, two without label); finally, a couple in my private collection, both from Hulata, 1.V1I1.1947 and 23-28.V1.1950. These, together with a male in the collection of Kibbutz Beit Ussishkin (Israel), taken at
Hula on 10.VI.1954, a series in the collection of Kibbutz Beit Gordon (Israel), taken at Tiberias, 28.V.1931, 17.V.1939, 3.V.1946, East of Bet-Yarah,
16.VI.1942 and Hula, 8.VI.1944, the male and females recorded by MORTON
(1924) from Hula, 24.V.1922 and the male recorded by SCHMIDT (1938) from
2 of - Fig. (a) Accessory genitalia Rhyothemis semihyalina syriaca, <5, lateral view; (b)
ultimate abdominal of of the segments a female same, showing the valvules.
3 Ferum near Rosh Pina, Hula Lake, 25.VII.1928, are the only existing specimens
that I know of, in all about thirty at the most.
I might add that the hyaline fringe in the hindwing is wider in females than in
males, and that the base of the hindwing is less widened in R. semihyalina syriaca than in R. semihyalina semihyalina (Fig. la, b). Selys’ ’’Syrian”
specimens might well have come from the Lake Hula area, since all other known
specimens were collected here. Structurally, the subspecies is identical with the
- Fig. 3 Urothemis edwardsi hulae subsp.n.: (a) Abdominal colour pattern of the male; (b)
the Accessory genitalia of male, lateral view; — (c) Appendices of the male, lateral view; —
(d) Ultimate abdominal segments ofa female, showing the valvules.
4 nomino-typical one in both sexes (Fig. 2a, b). The disjunction is considerable; typical semihyalina occurs in Algeria (SELYS, 1849) and further in the whole
Ethiopian region (PINHEY, 1962).
UROTHEMIS EDWARDSI HULAE SSP. NOV.
Figures 3 a-d, 4 a
Material. - Holotype: a fully adult, pruinose d, labeled Huleh (Hula), 9.VI. 1952,
deposited at the Department of Zoology, University of Tel Aviv. Paratypeseries:
7dd, Hula, 23.VI.1952, 699, Hula, 23.VI.1952, Department of Genetics, The Hebrew Uni-
versity of Jerusalem; Id, 19, same label, author’s collection; Id, 19, same labels, British
Museum (Natural History); Id, Hula, 4.XI.1946, author’s collection; 19, Jericho,
16.VII.1942 (leg. Bytinski-Salz), 399, Hulata, 1.VIII.1947 (leg. Bytinski-Salz), 2 66, Hula,
9.VI.1952 and 19, Ulmania (Hulavalley), 2.VII.1947, all at the Department of Zoology, Tel
Aviv University.
Fig. 4 (a) Hind wing of Urothemis edwardsi hulae subsp.n.; - (b) Hind wing of U. e.
edwardsi, Maun, Botswana (E.C.G. Pinhey leg.).
5 Additional material. - 19, Gonen, 5.VI.19S3, coll. Kibbutz Beit Ussishkin; a series in both sexes, coll. Kibbutz Beit-Gordon, from Dan, 1.V1I.1953, Gonen, 5.VI.1953, and Sede
Nehemya, without date. Further, a female was reported by Morton (1924) from Hula,
24.V.1922 and 4 66, 499 from Rosh Pina-Hula lake (”an Wassergraben”), 23.VII.1928, by
SCHMIDT (1938).
Male: structurally identical with U. e. edwardsi (Selys). Hamuli and appendices as in Figures 3b, c. Differs from nominal subspecies in the extent of the basal spot on the hindwings which is greatly reduced (Fig. 4a).
of Female: structurally identical to U. e. edwardsi (Selys). Structure val-
reduced male. vules as in Figure 3d. The basal spot on the hindwings as in the
Distribution: all female specimens, save a single taken near Jericho, were captured at lake Hula and surroundings. The nominal subspecies, although described from Algeria (SELYS, 1849), is widely distributed in tropical Africa
(P1NHEY, 1962).
DISCUSSION
One rarely has the privilege to deal with the fauna of a lake that has been famous since antiquity. As stated by COWGILL (1969), a first rough estimate of its superficy was given by Flavius Josephus, somewhere between 69 and 79 A.D.
Throughout history, stray notes on the lake were published by visitors to the
it holy land, some claiming that might completely dry up during summer, others
of and that surround it. giving descriptions its shape, depth, the papyrus swamps
It is the first lake on the course of the River Jordan, and therefore, the parts of
of it directly adjacent to the channel the river can never have been completely
much dry. The second and major lake, Lake Tiberias (Yam Kinneret) has a poorer dragonfly fauna than Hula, as have many large and deep lakes.
The events of interest to this paper took place, presumably some 35-30,000
of the Rift north of the Hula rather years ago, and are the damming Jordan by
of the basins of complicated volcanic activities, finally resulting in a separation
and transformationof the Jordan and the Litani Rivers, the a previously existing swamp into the Hula lake. While this process was taking place, the Wiirm glaci- ation was in evidence in Europe and a corresponding pluvial existed in North
Africa and Egypt. Although the synchronism of Pleistocene glaciations and pluvials is not rigorous (there have been more pluvials than glaciations), they
shown for were nicely simultaneous during the Wiirm, as BUTZER (1971) has
Libya and Egypt. For Israel, PICARD (1963) arrives at the conclusion that no
appreciable climatic changes occurred during the late Pleistocene, although DAN
(1961) gives pedological evidence that the basaltic soils of the upper Jordan
valley are typical of a climate with a precipitation of about twice the present.
This leads to the suggestion that the Arava valley and the Sinai were more humid
than Some to-day and were no barrier to non-migrant dragonflies. Ethiopian
6 that had followed the species Nile-pathway could thus cross the Sinai and Negev
deserts and reach the Jordan where valley, they moved North up to Hula, but apparently did not cross the Yarda basalt between the Naphtali mountains (of
Cretaceous age) and the Hermon (of Jurassic age). The cessation of this humid
occurred period in an oscillatory way (BUTZER, 1971), and may have become efficient around 15000 BP. It resulted in a progressive shrinking of the number of viable for in the South but also the biotopes dragonflies, not only in present
Syrian desert. This is proven by the distribution of e.g. Brachythemis fuscopal- liata which lives (Selys), now in the great marshes of Iraq, but has left relict populations in the Adana peninsula (S. Turkey) and at Hula (DUMONT, 1972).
the habitats suitable for the reduced the Likewise, Ethiopian immigrants were to upper Jordan valley, roughly the Hula lake and its surrounding marshes.
The for arguments placing these Ethiopian immigrants in the Wiirm(and not
at an earlier date) follow from:
(1) their absence North of the Jordan valley. The objection might be raised that we do not know enough about the fauna of The Lebanon and Western Syria to make such a statement acceptable. Indeed, two habitats exist that might have had the same role as lake Hula, viz. Lac de Homs (Syria) and Amik Golii (Hatay,
Turkey). I visited Amik Golii in July 1973, and found it completely drained and its former bed planted with cotton. Whatever dragonfly fauna existed here will forever remain unknown. Lac de Homs, which I did not see, was visited by
GADEAU DE KERV1LLE (1926), who appears to have been the only one to
collect here. collection dragonflies His contains none of the Hula endemics, but
rather small and it was a thorough study of this area would definitely be of interest.
(2) If one compares the calopterygid fauna of the Jordan and the Litani, one is struck by the presence, on the Jordan, of Calopteryx syriaca (Gene), and of both C. and C. syriaca (Gene) hyalina (Martin) on the Litani. The status of the latter is uncertain (full species? subspecies? a local conditionof syriaca?), but the fact which matters is that, at least, a genetic difference between populations North and South of the Yarda basalt exists, and, thus, that the barrier has been effective both sides. well on As is known, calopterygids are tightly associated with running water and cannot easily cross a mountain barrier.
CONCLUSIONS
The remarkable endemisation of dragonflies along the mediterranean coast of the Near East may well be understood bij the geological and climatic changes that took place since the end of the Tertiary epoch. BODENHEIMER (1938) believes that that, at time, a steppe fauna of Irano-Turanian origin extended through the Near-East to North Africa and the steppes of South Spain. As explained elsewhere (DUMONT, in preparation), the present-day dragonfly
7 fauna of Turkey shows that, in fact, at the end of the Tertiary, the Odonata of the Eastern Mediterranean (and of a large part of Europe as well) had a pro- nounced Oriental (i.e. not only Central-Asiatic) facies. Remnants of this fauna, either primitive species or species and whole genera that evolved from these, are still in evidence here. But in the course of the Pleistocene, owing to the wax and wane of precipitations, immigrations from the African continent took place.
There has certainly been more than one Ethiopian vague, and e.g. Pseudagrion
has reached the much earlier syriacum (Selys) Near East at a date than the
Hula-fauna (DUMONT, 1974). Proof is that this animal has reached full specific
status (it is remotely allied to P. kersteni Gerstaecker, an African species that
does not even occur in Egypt any more) and has a much wider distribution,
including The Lebanon and Lac de Homs (GADEAU DE KERV1LLE, 1926). If
one wishes to gain insight into the recent history of the West-Palearctic fauna,
the Near East is the place where such knowledge should be gathered. Para-
HUTCHINSON phrasing (1965), it is a theatre where evolution is being played.
Evolution linear function of and stimulated is not a time, seems to be greatly by fluctuating environmental conditions. One might find great intellectual pleasure in speculating about the synchronism in the development of the Hula fauna and the important step for humanity taken by the tribes at Jericho, switching from nomadism to agriculture, another probable consequence of an environment that became more arid.
EPILOGUE
It is a bit sad to have to add that two of the endemics of Hula are probably no
more in existence. Lake Hula, situated in a fertile plain was at the same time a
centre for the propagation of malaria. Therefore, it was drained between 1951
of and 1958, and only part of the papyrus swamp and some the open water was
but the preserved. The fatal thing that happened was probably not the drainage,
subsequent cultural eutrophication which resulted in the appearance of heavy water-blooms due to blue-green algae. As a consequence, the dragonfly fauna
but was greatly altered. Not only Brachythemis fuscopalliata (DUMONT, 1972),
also Rhyothemis semihyalina syriaca and Urothemis edwardsi hulae disappeared
at this occasion. Although the supervisor of the natural reserve (A. Freidberg, in
material litt.) claims to still see the former species now and then, there is no
evidence for this. The last captures took place in 1954, i.e. while the drainage
around be was proceeding. Both must still have been very common 1953, as can
Prof. J. Wahrman in June of judged from a random sample taken by (Jerusalem)
doubtful to be the first which that year. Therefore, this paper has the merit in a
after become new dragonfly taxon ( U.e. hulae) is described it has extinct.
8 ACKNOWLEDGEMENTS
I am indebted to Prof. J. KUGLER and Dr. A, FREIDBERG (Tel Aviv) and to Prof. J.
WAHRMAN (Jerusalem) for making available the dragonfly collections entrusted to them.
I To Dr. E.C.G. PINHEY (Bulawayo), am obliged for sending material for comparison from
the Ethiopian region.
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